A study on herbivores of the red alga Mazzaella splendens in Oregon also includes grazing activity of the black-leather chiton Katharina tunicata.  Removal of all Katharina from intertidal rocky benches leads to re-establishment of Mazzaella in areas where it was previously absent, and removal of limpets Lottia pelta along with Katharina accentuates this.  Removal of Katharina has a strong positive effect on density of limpets, suggesting competitive interactions between them.  Gaines 1985 Ecology 66: 473.

NOTE formerly Iridaea cordata

A black leather chiton Katharina tunicata in an area
of kelp Hedophyllum sessile and coralline algae.
Some limpets are present, at least one of which
is Lottia pelta (in the 2 o'clock position) 0.25X

There have been several studies on effects of chiton herbivory on mediating structure of west-coast intertidal communites.  One of these, conducted in San Juan Island, Washington, concerns the role of herbivory by leather chitons Katharina tunicata¹ in affecting competitive interactions between marine plants².  The authors create 3 types of treatment plots: chiton-removal³, CONTROL, and chiton-addition, and monitor abundance and diversity of the various plants over a 4-yr period.  In the areas where Katharina are removed, algal diversity and abundance increase, leading at the end to a multi-story intertidal kelp bed.  CONTROL sites show year-to-year fluctuations in abundance of the large food-alga Hedophyllum sessile, but otherwise densities remain more-or-less unchanged. 

Effects on several "corticated macrophytes" are shown graphically on the Left and in before-and-after photographs on the Right for a chiton-removal area.  In the Katharina-addition areas, abundance and diversity of algae decline (data not shown here).  In fact, the only plants to thrive under the intense grazing pressure in the chiton-addition areas are surfgrasses Phyllospadix scouleri and diatoms, the former thought possibly to be indigestible by Katharina. Later removal of large kelps from chiton-removal and CONTROL areas lead to rapid increase in abundance and diversity of other algae.  The kelps apparently suppress the growth of smaller forms beneath them.  Because these effects are most pronounced in areas lacking both chitons and kelps, the authors suggest that herbivory and competition have negative, additive effects on the smaller algae.  Duggins & Dethier 1985 Oecologia 67: 183.

NOTE¹  in the areas of San Juan Island selected for study, Katharina is the dominant herbivore, reaching densities of up to 100 ^. m^-2 and sizes of 12cm length

NOTE² other plants, including crusts, diatoms, and surfgrass, are also monitored, but the data are less interesting than those for macroalgae and thus are not included here. The data set presented here represents "corticated macrophytes": Gigartina spp., Fucus spp., Rhodomela larix, Odonthalia spp., Botryoglossum sp., Plocamium sp., Iridaea spp., and other unidentified red species

NOTE³ chitons that wander into the removal areas are cleared out every 4-5mo.  This housekeeping maintains densities in the removal areas at about 8% of original, and in the addition area atabout 180% of original

A later study by the same authors compares the effects of removing black-leather chitons Katharina tunicata from low intertidal regions in San Juan Island, Washington and Torch Bay, Alaska over a 3-yr period.  Interestingly, while this yields dramatic changes in community structure in San Juan Island, Washington (see histogram on Left), little effect is seen in Alaska. In the Washington experiment the absence of K. tunicata for 3yr results in a kelp bed developing in the low intertidal region consisting of Hedophyllum sessile, Alaria marginata, and Nereocystis luetkeana, and it persists. Smaller algae later increase in abundance and diversity and, after 3yr, a multi-layered algal canopy is present. 

In comparison, effects of removal of chitons on kelps and other algae in Alaska are variable and not related to the presence or absence of Katharina. Note in the graph that while kelps increase in % cover in Alaska following removal of Katharina, they do so in both removal and control plots.  The authors caution that the presence of a strong interaction in one area does not guarantee its presence in another, even if community composition is similar in the 2 regions.  Dethier & Duggins 1988 Mar Ecol Progr Ser 50: 97.

NOTE  chiton densities are similar in the two areas, ranging from 21-57 individuals ^. m^-2

NOTE  replicate sites are cleared in each geographical area, but only results for one replicate are shown here

In Barkley Sound, British Columbia, black-leather chitons Katharina tunicata inhabit the same intertidal areas as does a potential food and shelter species, the kelp Hedophyllum sessile.

By manipulating densities of Katharina in 3 types of experimental plots¹, namely, low, CONTROL, and high densities², and comparing survivorship of Hedophyllum in each, the researchers find that Katharina affects survival of both juvenile³ (<1cm holdfast diameter) and young adult (1-4cm) plants, but not of adult (>4cm) plants. Note in the histogram that Hedophyllum densities decline in all treatment groups, including the CONTROLS over the 1-yr study period, in major part owing to winter-storm damage.  However, by April 1996 there is a significant difference between densities in the low and high treatment areas, and after this date there are no surviving Hedophyllum individuals in the high-density treatment areas. The effect on juvenile kelp plants is direct, in that the chitons eat them up. Effects on young adult plants, however, are indirect, in that the chitons damage the holdfasts resulting in greater losses to waves.  Apparently, the chitons seek out protection from desiccation or waves by burrowing into and edging under the holdfasts, thus loosening their attachment to the rock. The findings of the study provide evidence for a refuge in size for Hedophyllum from one of its principal grazers.  Markel & DeWreede 1998 Mar Ecol Progr Ser 166: 151. Photographs courtesy Russ Markel, University of British Columbia & Bamfield Marine Sciences Centre.

NOTE¹ the authors do not use fence barriers to delineate these plots; rather, they employ natural topographical features such as surge channels and outcroppings to restrict the chitons’ movements

NOTE² experimental densities are based on the original average density of Katharina in the 18 treatment areas, adjusted to 25% of this value in the “low-density” treatments and to 75% in the “high-density” treatments. There are 6 replicates of each of the 3 treatments

NOTE³ algal sporelings are highly vulnerable to grazing invertebrate herbivores and this early phase of life has been termed a “herbivory bottleneck” by marine algologists.  Although the same principles applies to larvae of marine invertebrates, there seems to be no comparable “carnivory bottleneck” referred to in the literature

In San Juan Island, Washington the chiton Katharina tunicata  associates closely with the canopy-forming brown alga Hedophyllum sessile¹.  In this region the alga’s canopy cover changes from about 60% in winter to 90% in summer.  Experiments² involving addition of bird-exclusion and shading screens³ show that the association owes to selection of shaded microhabitats by the chiton, and not to protection from predators, such as seagulls, or presence of a potential food.  The author notes that sea stars Pisaster ochraceus represent the chief predators of Katharina in cageless control plots and in shade-treatment cages, but not in the enclosed bird-exclusion cages, which are inaccessible to thesea stars.  In summer, 33% of Pisaster’s diet is made up of Katharina and, in winter, 17%. 

Data on seasonal changes in levels of heat-shock proteins (Hsp70⁴, see graph on Right) indicate that the chitons experience sublethal temperature stresses during spring and summer. Note in particular that the seasonal onset of daytime low tides occurring in mid- to late-spring is associated with increased levels of Hsp70, indicating that this may be a time of significant physiological stress for the chitons.  During these stressful periods Katharina’s intertidal distribution appears to be limited by the availability of shady refugia.  Note in the body-temperature graph (above Left) the marked effect that shade, whether naturally created by Hedophyllum or within the “artificial"-shade cages, has on mitigating temperature effects in Katharina. Burnaford 2004 Ecology 85: 2837.

NOTE¹ this canopy-forming perennial kelp delineates a specific level in the low intertidal zone of rocky beaches.  So important is it in community dynamics of the shore that it has been termed a  “foundation species” and an  “ecologically dominant species” by various authors

NOTE² the data presentation given in the paper on numbers of chitons within each treatment over the nearly 3-yr study period is complex and is not presented here

NOTE³ these are upside-down vinyl-coated wire-mesh baskets, fastened to the rock surface.  Other top-less baskets, presumably accessible to birds, serve as control treatments.  Baskets with vexar-mesh tops represent shade treatments to mimic the natural canopy-shading of Hedophyllum.  Holes are cut in the sides of all cages at rock level to allow the chitons ingress and egress

NOTE⁴ the expression of Heat Shock Proteins (HSPs) is commonly used to assess stress in animals resulting from physical factors such as temperature, UV light, and other stresses.  HSP70 is actually a complex of these proteins but, for convenience, is expressed collectively as “Hsp70” by the authors.  More on HSPs in the ODYSSEY can be found at LEARN ABOUT ABALONES & RELATIVES: PHYSIOLOGICAL ECOLOGY: HEAT-SHOCK PROTEINS, and LEARN ABOUT MUSSELS: LIFE IN THE INTERTIDAL ZONE: HEAT-SHOCK PROTEINS

The gumboot chiton Cryptochiton stelleri is the largest chiton in the world, yet little is known of its population dynamics or ecology.  An investigation of C. stelleri’s distribution at 6 rocky intertidal sites by a researcher at the Oregon Institute of Marine Biology, Charleston helps to redress the situation.  Results show highly patched and clumped distributions, mostly within wave-protected coves.  The preference for wave-protected areas may in part owe to a weaker attachment tenacity than other wave-tolerant chiton species (e.g., black leather chitons Katharina tunicata), but probably more to a greater abundance in such habitats of thin-bladed algal species such as Mazzaella splendens and Cryptopleura sp. favoured as food by C. stelleri.  Age-frequency plots reveal that recruitment is sporadic, low, and site-specific (see representative plot below for Cape Blanco).  Juveniles less than 15cm in length are rare, and when present tend to be found within pits in sandstone created by purple sea urchins.  The author suggests that the slower, swirling water-conditions in cove habitats may favour the trapping and settlement of larvae¹, and later survival of recruits. The age-frequency data presented in the histogram are based on growth lines in shell plates, and indicate that maximum age is about 40yr.  Results from monitoring of tagged² individuals show that C. stelleri are homebodies, and move less than about 20m on average over a 6mo period.  The author notes a lack of evidence of age-related movements of individuals within the vertical range of intertidal zone.  Laboratory experiments in which 10-30cm-sized individuals are maintained together with sea-stars and crabs in aquariums for 2mo reveal no predation³.  The author suggests that population-connectivity between sites on the Oregon coast is low, but notes that the necessary genetic work has not yet been done to show this.  The study is a welcome contribution to an under-investigated group of intertidal herbivores.  Lord 2011 Malacologia 54 (1-2): 147. Photograph courtesy Linda Schroeder, Pacific Northwest Shell Club, Seattle, Washington.

NOTE¹  larvae are lecithotrophic and are able to settle within a few days of hatching

NOTE²  individuals are tagged by inserting plastic zip ties through the mantle edge bearing numbered fish tags.  Tag retention is poor, with only 14% being found after 6mo

NOTE³  predatory species tested include sea stars Evasterias troschelii, Pisaster ochraceus, and Pycnopodia helianthoides, and crabs Pugettia producta and Cancer productus

Gumboot chiton Cryptochiton stelleri
partially submerged in a tidepool 0.5X